Redalyc.Record of the Rare Oceanic Salp Helicosalpa Komaii

Revista Mexicana de Biodiversidad ISSN: 1870-3453 [email protected] Universidad Nacional Autónoma de México México

Hereu, Clara M.; Suárez-Morales, Eduardo; Lavaniegos, Bertha E. Record of the rare oceanic salp Helicosalpa komaii (Tunicata: Thaliacea: Salpida) in the Northeast Pacific Revista Mexicana de Biodiversidad, vol. 85, núm. 2, 2014, pp. 624-629 Universidad Nacional Autónoma de México Distrito Federal, México

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Research note

Record of the rare oceanic salp Helicosalpa komaii (Tunicata: Thaliacea: Salpida) in the Northeast Pacific

Registro de una especie oceánica rara de salpa Helicosalpa komaii (Tunicata: Thaliacea: Salpida) en el Pacífico nororiental

Clara M. Hereu1 , Eduardo Suárez-Morales2 and Bertha E. Lavaniegos3

1Facultad de Ciencias, Universidad Autónoma de Baja California. Carretera Tijuana-Ensenada, Km 103, 22860 Ensenada, Baja California, Mexico. 2Departamento de Ecología y Sistemática Acuática, El Colegio de la Frontera Sur. Av. Centenario, Km 5.5, 77014 Chetumal, Quintana Roo, Mexico. 3Departamento de Oceanografía Biológica, Centro de Investigación y de Educación Superior de Ensenada. Carretera Tijuana-Ensenada Km 107, 22860 Ensenada, Baja California, Mexico. [email protected]

Abstract. We report for the first time the presence of a rare salp Helicosalpa komaii (Ihle and Ihle-Landenberg, 1936) on the west coast of Baja California peninsula (26°52.7’ N, 117°09.2’ W). The single specimen recorded was a solitary zooid of 200 mm length and 90 ml of displacement volume. It was recognized by having a distinctive highly convoluted heart-shaped dorsal tubercle. The total number of muscle fibers in the solitary zooid (MI to MVII= 517) surpassed that of the other 2 known congeners supporting its identity as H. komaii. H. komaii increases to 29 the total number of salps species reported in coastal and oceanic waters off north-central Baja California (gamma diversity). We hypothesize that the presence of H. komaii in the study area may be the result of onshore intrusions of subtropical waters originating west-southwest of the California Current domain.

Key words: salps, marine zooplankton, pelagic tunicates, California Current, Baja California.

Resumen. Se registra por primera vez la presencia de una especie rara de salpa Helicosalpa komaii (Ihle y Ihle Landenberg, 1936) en la costa oeste de la península de Baja California (26°52.7’ N, 117°09.2’ O). El ejemplar registrado fue un zooide solitario de 200 mm longitud total y una biomasa de 90 ml (volumen desplazado). El ejemplar fue reconocido por su característico tubérculo dorsal en forma de corazón. El número total de fibras musculares en el zooide solitario (MI a MVII= 517) superó el de los otros dos congéneres conocidos, apoyando su identificación como H. komaii. El registro de H. komaii incrementó a 29 el número total de especies de salpas registradas en aguas costeras y oceánicas del centro-norte de Baja California (diversidad gamma). Nuestra hipótesis es que la presencia de H. komaii en el área de estudio se debe a su acarreo hacia la costa por agua subtropical procedente del oeste-suroeste del dominio de la corriente de California.

Palabras clave: salpas, zooplancton marino, tunicados pelágicos, corriente de California, Baja California.

Salps are pelagic tunicates widely distributed in all to the worldwide 44 known species (Van Soest, 1998; oceans, comprising the most diverse order among the Govindarajan et al., 2010). thaliaceans (Van Soest, 1998). The order Salpida contains Cyclosalpinae salps of the genus Helicosalpa (Todaro, 1 family with 2 subfamilies: Cyclosalpinae Yount, 1954 1902) are among the most rarely collected. Currently, including 2 genera, and Salpinae Yount, 1954 with eleven the genus contains 3 species: Helicosalpa virgula (Vogt, genera. Salps have wide zoogeographic distribution and 1854), H. younti Kashkina 1973, and H. komaii (Ihle and the discovery of new species is considerably rare. Only Ihle-Landenberg, 1936), a species originally described a few new species have been added in the last decades as Cyclosalpa komaii (Van Soest, 2013). Helicosalpa virgula has the widest zoogeographic distribution and is the most frequent species reported in the genus. This Recibido: 30 marzo 2013; aceptado: 22 enero 2014 large species has been recorded from different localities Revista Mexicana de Biodiversidad 85: 624-629, 2014 DOI: 10.7550/rmb.36638 625

in the Mediterranean, the Atlantic Ocean and also from 120.70 (26°52.7’ N, 117°9.2’ W; 17.19 °C and 33.44 PSU the Indian and the Pacific Oceans (Fig. 1). Fewer H. at 10 m depth) of the research program “Investigaciones younti records exist, known only from the central Pacific, Mexicanas de la Corriente de California” (IMECOCAL, off India (Van Soest, 1974), and from Chilean waters; Fig. 1) (Lavaniegos et al., 2006). The zooplankton sample its aggregate zooid was recently described by Esnal et was collected with a Bongo net (71 cm mouth diameter, 0.5 al. (1998). Helicosalpa komaii is the rarest species with mm mesh size) obliquely towed from 200 m depth to the only 1 solitary and 30 aggregates zooids recorded in the surface. The zooplankton sample was preserved in a 4% western Pacific (Komai, 1932), a single solitary zooid formaldehyde solution buffered with sodium borate. The reported in Central Pacific waters (Yount, 1954), a chain The specimen of H. komaii was deposited in the collection of 8 aggregates in the Gulf of California (Madin, 1968) and of zooplankton at El Colegio de la Frontera Sur, Unidad 14 aggregates zooids collected in the Indian Ocean (Van Chetumal, in Chetumal, Quintana Roo, Mexico (specimen Soest, 1974). There are 2 recent additional records of the catalog number ECO-CH-Z-6300). species, from southwest Taiwan (Tew and Lo, 2005) and The solitary zooid specimen of H. komaii collected off the coast of Peru (Ayón et al., 2008). Unfortunately, and examined was physically damaged, since a section of neither figures nor zooid specifications were provided by the outer gelatinous tunic from the body wall was partially these authors. Here we report the extension range of H. broken off. However, a careful handling of the organism komaii in the Northeast Pacific, particularly off the coast in a wide glass container allowed a detailed observation of Baja California peninsula, Mexico. of the main taxonomic features that distinguishes the The zooplankton sample containing the specimen of solitary form of H. komaii from the other species known Helicosalpa komaii reported here was obtained during a of the same genus: H. virgula and H. younti. Distinctive nighttime trawl carried out on February 14, 2004 at station H. komaii taxonomic features are: the presence of 2 ventral

Figure 1. Worldwide records of the genus Helicosalpa: H. komaii (squares), H. younti (diamonds) and H. virgula (circles). The sampling grid off Baja California by IMECOCAL Program is detailed in lower left corner showing northern (N) and southern (S) sectors and the location of records of H. komaii off Baja California and in the Gulf of California (GC). Locations were obtained from: Thompson, 1948; Yount, 1954; Madin, 1968; Hubbard and Pearcy, 1971; Van Soest, 1974; Blackburn, 1979; Madin et al., 1996; Esnal et al., 1998; Tew and Lo, 2005; Hereu et al., 2006; Wiebe et al., 2006; Ayón et al., 2008; Weikert and Godeaux, 2008; and Lavaniegos (unpublished data). Average Sea Surface Temperature taken from WOA09 at NOAA´s “Ocean Climate Laboratory” free access server (Locarnini et al., 2010). 626 Hereu et al.- New record of Helicosalpa komaii in Northeast Pacific

longitudinal muscles (found only in Helicosalpa, absent in Cyclosalpa) stretching between the intermediate muscles and muscle VII, 1 dorsal longitudinal muscle, and a highly convoluted heart-shaped dorsal tubercle (Figs. 2, 3). Another specimen corresponding to the aggregate form of the genus Helicosalpa was also found in the same zooplankton sample, bearing a horn-like antero-dorsal projection, which is also present in H. virgula and H. younti. The 8 mm length aggregate zooid resembled the sinistral individual of H. komaii (figure 6B in Komai, 1932) in the presence of a roundish postero-ventral protrusion (not tapering posteriorly into a narrow point). The dorsal tubercle was not as convoluted as described by Van Soest (1974) for the aggregate zooid of H. komaii. However, Esnal et al. (1998) mention that this character is variable with size, being more convoluted in larger individuals. Figure 3. Heart-shaped dorsal tubercle of the solitary zooid of The disposition of body muscles and number of muscles Helicosalpa komaii from off Baja California. fibers were difficult to observe in this small aggregate specimen, therefore its definitive identity remains dubious and emphasis is given on the collected solitary specimen. (94 mm) collected south of Hawaii, outlining only slight Only 2 other studies provide biometric measurements differences between his and Komai´s specimen. According of the solitary form of H. komaii. Komai (1932) to Nakamura and Yount (1958), this species is among described the largest specimen known (230 mm) from the largest salps known in the Pacific Ocean, together Seto, Japan, and Yount (1954) described another solitary with Tethys vagina and Salpa maxima. We compared our specimen based on the taxonomic characters used in the 2 previous morphometric descriptions, particularly taking care of the muscle arrangement and the presence/absence of other relevant taxonomic characters. Solitary form. The solitary specimen from Baja California was approximately 200 mm length and 60 mm height, with a biovolume of 90 mL with 6 years of preservation after collection. Body muscles with variable number of muscles fibers (from 50 to 90). Total number of muscles fibers from M I to M VII= 517. Muscles I and II conjoined near mid-dorsal line and continuing as 1 dorsal longitudinal muscle that joins M VII; it stretches backward ventrally at both sides of the body to form the paired ventral longitudinal muscles. Muscle VIII, named muscle Y in Yount (1954) description, emerges from the union of ventral longitudinal muscle with MVII, continuing dorsally as a single broad muscle (i.e., non-branched as in Komai´s description). Body muscles III-VI free dorsally, but fused ventrally to longitudinal muscles (Fig. 2). Oral musculature was similar to description in Yount (1954), but there is a muscle joining the posterior dorsal lip with retractor (intermediate) muscle before the latter fuses to M I dorsally, as described in Komai (1932). A conspicuous Figure 2. Schematic representation of general morphology and heart-shaped dorsal tubercle highly convoluted (Fig. 3). muscle band distribution of the solitary zooid of Helicosalpa This was the main morphological feature to identify H. komaii: A, lateral view; B, dorsal view. Muscle numbers in Roman komaii at first sight. The alimentary canal had generic (I to VIII) from anterior (oral) to posterior (atrial) position. Oral characteristics, extending obliquely and dorsally to gill-bar, musculature reconstructed following Yount (1954). bending posteriorly at level of M I. Two prominent caeca Revista Mexicana de Biodiversidad 85: 624-629, 2014 DOI: 10.7550/rmb.36638 627

are present at the opposite extreme of the intestine. A stolon have been useful indicators of intrusions of warm oceanic is formed between muscles VI and VII and extends forward waters into colder zones (Fraser, 1962; Blackburn, 1979; in the mid-ventral line, reaching the anterior extreme of the McAlice, 1986; Sims, 1996; Iguchi and Kidokoro, 2006). salp. It was not possible to determine the place where the A recent study on the variability of different zooplankton stolon emerged to the body surface because of loosening groups’ abundance off Baja California within the period of the tunic in this section of the body. Strobilation was 1998-2007 denoted a marked seasonality in several taxa evident along the stolon; however, the forming aggregates including salps. This seasonality is more notorious at the were small and their arrangement pattern along the stolon southern eco-region (Fig. 1) due to a stronger influence of was more irregular than the one typically observed in H. a tropical oceanic influx in that sector of the IMECOCAL virgula or other salp species. Atrial languets were absent, area (Durazo et al., 2010; Lavaniegos et al., 2010). This similar to the specimen described in Yount (1954), but tropical influence is more pronounced during El Niño this is an ambiguous morphological feature because atrial warm years and it is reflected in shifts in the species languets were present in the specimen described by Komai arrangement and geographic range extensions (Jiménez- (1932). Pérez and Lavaniegos, 2004; Linacre, 2005; Lavaniegos According to previous descriptions, the species is and Ambriz-Arreola, 2012). The winter 2004, when characterized by the presence of luminous organs forming the specimen of Helicosalpa komaii was collected, was a continuous line on each side of the body. This feature was followed by the moderate 2002-2003 El Niño that resulted not observed in the Baja California collected specimen, but in a zooplankton community rich in tropical species off vestiges of the luminous organs were detected. Overall, Baja California (Lavaniegos and Ambriz-Arreola, 2012). these structures are not as conspicuous in H. komaii as they Helicosalpa komaii is distributed in low and mid latitudes are in the other 2 species of the genus. Yount (1954) noted in the Pacific Ocean, therefore its occurrence in the that they apparently are formed by sparse masses of cells. offshore station in the southern Baja California waters may Chemical preservation and time elapsed after collection have resulted from intrusion of warm water from the west- may distort the appearance of this structure (Esnal et al., southwest toward the coast. Similarly, the presence of H. 1998). virgula and other warm water salp species (e.g. Thetys The total number of muscle fibers (M I-M VII) appears vagina, Ritteriella amboinensis, Thalia rhomboides) off to be a taxonomic significant character to distinguish H. Oregon (Hubbard and Pearcy, 1971) and Baja California komaii from the other 2 species of the genus. This character (Berner, 1967; Hereu et al., 2006) were associated to the is size and age independent and remains practically onshore intrusion of tropical waters of west-southwest unaffected by chemical preservation (Esnal et al., 1998). origin and the weakening of the cold California Current Latitudinal variation in the total number of muscle fibers during anomalous El Niño years. The high abundance of within a species has been reported, with an increasing salps and other pelagic tunicates (i.e, doliolids) in the winter number of muscle fibers from tropical to higher latitudes of 2004, mainly in the southern eco-region, reinforces the (Van Soest, 1975). The high number of muscles fibers idea of a stronger influx of tropical oceanic waters in determined for our specimen (517) surpasses the range that area (Lavaniegos et al., 2006; Durazo, 2009). The known for H. younti (with 343 and 346 determined in determination of salp taxonomic identity in the remaining 2 solitary zooids) (Esnal et al., 1998). It is also 2-fold stations off Baja California in the winter 2004 cruise will higher than that reported in H. virgula whose solitary help to reinforce the hypothesis. zooids have on average 264 and 218 total muscles fibers This is the first report of Helicosalpa komaii in the in cold and warm waters, respectively (Van Soest, 1975). California Current System (CCS) and the second report in Unfortunately, information on the total number of muscle the Northeast Pacific. With this new record, the number of fibers was not provided in the 2 previous descriptions salp species (gamma diversity) detected in Baja California of the solitary form of H. komaii and no intra-species waters is increased to 29 and to 30 in the Mexican Pacific comparison of this character is possible. We propose that (Berner, 1967; Esnal, 1976; Hereu et al., 2006; Lavaniegos this is another significant taxonomic character to identify and Hereu, 2009; Hereu et al. 2010). This report provides the solitary specimen of H. komaii. For the aggregate zooid valuable information regarding the distribution of salps in of the species, the average total number of muscle fibers the CCS given that there are no historical records of H. from MI to MIV is 90 (range= 81-97), doubling that of H. komaii previously identified in the area, including the long virgula aggregates (range= 27-51; Van Soest, 1974) and term CALCOFI sampling program of more than 50 years similar to the upper limit of the range (65-90) reported for (Berner, 1967; Blackburn, 1979; Lavaniegos and Ohman, H. younti (Esnal et al., 1998). 2003). Ecosystem changes were detected within the CCS Salp biogeography is relevant because several species from changes in the assemblages of pelagic tunicates 628 Hereu et al.- New record of Helicosalpa komaii in Northeast Pacific

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